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A review of dipterocarps - Center for International Forestry Research

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Biogeography and Evolutionary Systematics <strong>of</strong> Dipterocarpaceae<br />

the herbarium collections <strong>of</strong> Asia (Forest <strong>Research</strong><br />

Institute Malaysia, Kepong including Symington’s<br />

collection; Bogor in Indonesia, Peradeniya in Sri Lanka,<br />

Bangkok in Thailand) and Europe (Kew in U.K, Paris and<br />

Lyon in France, Leiden in Netherlands) where large<br />

collections including Ashton’s, Meijer’s, and Maury’s,<br />

are preserved.<br />

There is a certain complementarity in the above<br />

works, but a synthetic classification integrating all<br />

previous results is still not available. Traditional and<br />

modern approaches will have to be integrated, including<br />

DNA and mathematical analyses, as well as the use <strong>of</strong><br />

computer systems <strong>for</strong> determination and treatment <strong>of</strong><br />

the data.<br />

Main aspects <strong>of</strong> Maury-Lechon’s classification (see<br />

also Tables 1, 2, 8)<br />

The taxonomical levels have intentionally been left<br />

without <strong>for</strong>mal definition to serve as a base <strong>for</strong> further<br />

research. The relative position <strong>of</strong> these levels is much<br />

more important than their names. However, to facilitate<br />

the explanations, the more proximal names usually<br />

adopted <strong>for</strong> these divisions were used (Maury-Lechon<br />

1979a).<br />

The separation <strong>of</strong> Monotoid taxa from the<br />

Dipterocarpaceae underlines the differences that<br />

introduce heterogeneity when these taxa are put together<br />

with the Asian group in the same family. The grouping <strong>of</strong><br />

Monotaceae and Dipterocarpaceae in a supra-familial<br />

joint division (order or suborder), reminds the greater<br />

affinities <strong>of</strong> these two families among the other<br />

angiosperms. All the other groups aim to underline the<br />

closer resemblances on the basis <strong>of</strong> living biological,<br />

morphological and anatomical characters <strong>of</strong> the<br />

successive ontogenic phases (mainly fruit-seed, embryo,<br />

seedling) and the pollen types and structures.<br />

The characters <strong>of</strong> embryo-seedlings and pollens<br />

strongly emphasise the particular position <strong>of</strong><br />

Anthoshorea close to Doona and partly to Pentacme<br />

within the Anthoshorinae group, and their position near<br />

Hopea and Neobalanocarpus in the Imbricate group.<br />

Still stronger relations exist with the intermediary<br />

genus Dryobalanops leading directly to the Valvate<br />

group. Tighter connections <strong>of</strong> the latter genus occur<br />

within the Dipterocarpae subgroup (Dipterocarpus and<br />

Anisoptera) as well as with the Vaticae subgroup through<br />

Sunaptinae taxa first (Cotylelobium mainly, Sunaptea<br />

too), and then to Stemonoporus. Through Dryobalanops<br />

Taxonomical levels<br />

Supra-family<br />

level (1) (order<br />

or sub-order):<br />

Dipterocarpales (2 families)<br />

Family level Monotaceae (3 genera:<br />

(2):<br />

Pakaraimaea, Marquesia,<br />

Monotes)<br />

Dipterocarpaceae (2 infra-family<br />

groups: sub-families, 4 sub-groups:<br />

tribes, 9 sub-subgroups: sub-tribes,<br />

19 genera)<br />

Sub-family Imbricate [2 tribes (a & b), 3 sub-<br />

level (2):<br />

tribes, 9 genera, 19 sections]<br />

Valvate [2 tribes (c & d), 7 subtribes,<br />

10 genera, 8 sections]<br />

Tribe level (4): a) Hopeae (2 genera, 4 sections):<br />

Hopea (4 sections),<br />

Neobalanocarpus<br />

b) Shoreae (3 sub-tribes, 7 genera, 15<br />

sections):<br />

Anthoshorinae (3 genera): Doona,<br />

Pentacme, Anthoshorea (2<br />

sections)<br />

Shorinae (3 genera): Shorea (2<br />

sections), Richetia (2 sections),<br />

Rubroshorea (7 sections)<br />

Parashorinae (1 genera):<br />

Parashorea (2 sections)<br />

c) Dipterocarpae (2 sub-tribes, 3<br />

genera, 2 sections):<br />

Dryobalanoinae (1 genus):<br />

Dryobalanops<br />

Dipterocarpinae (2 genera):<br />

Dipterocarpus, Anisoptera (2<br />

sections)<br />

d) Vaticae (5 sub-tribes, 7 genera, 6<br />

sections):<br />

Upuninae (1 genera): Upuna<br />

Sunaptinae (2 genera):<br />

Cotylelobium, Sunaptea (2 sections)<br />

Vaticinae (1 genus): Vatica pro<br />

parte (2 sections)<br />

Stemonoporinae (1 genera):<br />

Stemonoporus (2 sub-groups)<br />

Vaterinae (2 genera): Vateria,<br />

Vateriopsis<br />

31<br />

another line <strong>of</strong> similarities leads to Anisoptera, Vateria<br />

and Vateriopsis.<br />

Pollen, embryo and seedling characters, as well as<br />

parsimony analysis <strong>of</strong> molecular data (Tsumura et al.<br />

1993, Wickneswari 1993), show similar conclusions. By<br />

the pollen surface Anthoshorea resemble<br />

Dryobalanops, Doona, Neobalanocarpus heimii and<br />

Cotylelobium, and a basic similarity exists between<br />

Dryobalanops and Dipterocarpus, as with cotyledonary<br />

shapes.

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